Chromosome numbers in species of
& Osman Beyazo
Hayırlıoâlu-Ayaz, S. & Beyazoâlu, O., Department of Biology, Faculty of Science,
Kara-deniz Technical University, 61080 Trabzon, Turkey Received 29 June 1999, accepted 2 February 2000
Hayırlıoâlu-Ayaz, S. & Beyazoâlu, O. 2000: Chromosome numbers in species of
Al-chemilla ser. Elatae (Rosaceae)in Turkey. — Ann. Bot. Fennici 37: 173–182. This paper presents the results of a karyological analysis of 17 species of Alchemilla L. sect. Alchemilla subsect. Calycanthum Rothm. ser. Elatae Rothm. The studies were carried out on plants from northeast Anatolia (Turkey). About 75% of the species have chromosome numbers 2n = 86 to 123. A few species have a lower number. The species are polyploids like the majority of species of sect. Alchemilla studied so far from vari-ous geographic areas.
Key words: Alchemilla, chromosome number, ﬂora of Turkey, Rosaceae
The genus Alchemilla L. of the family Rosaceae consists of perennial herbs with a woody rhizome and includes more than 1 000 species. The spe-cies have a mainly holarctic distribution and are distributed at various elevations in almost all types of habitats; most frequently they occur in moun-tains, in subalpine and alpine zones (Izmailow 1982).
In Turkey, the genus is, according to Pawlow-ski and Walters (1972), represented by 3 subsec-tions and 6 series belonging to sect. Alchemilla. Most of them are found in northeast Anatolia.
Alchemilla, with its numerous and variable forms, is a taxonomic challenge and it has been the subject for taxonomic studies since the end of the 19th century. Buser (1894), in his revision of
the Swiss taxa, applied a very narrow species con-cept and described numerous forms of Alchemilla, treating them as distinct species. The taxonomic tradition initiated by Buser was continued by bota-nists such as Juzepczuk (1941), Walters (1952), Rothmaler (1944), and Pawlowski (1972).
Cytological data for Alchemilla are very scarce. The papers by Turesson (1957), Wegener (1967), Izmailow(1981, 1982), and Hayırlıoâlu
and Beyazoâlu (1997a, 1997b, 1997c) are
particu-larly important;Turesson (1957) determined the chromosome numbers for 19 species, Wegener (1967) for 56, Izmailow (1981, 1982) for 11, and Hayırlıoâlu and Beyazoâlu (1997a, 1997b, 1997c)
From a cytological as well as taxonomic point of view, Alchemilla presents a difﬁcult subject. Izmailow (1981, 1982) reported that all studied
species collected from the Western Carpathians (Poland) are high polyploids, and in spite of the small size of the chromosomes, it is difﬁcult to get them sufﬁciently to spread and to count their numbers exactly. Diploid cytotypes are as yet un-known.
Previously, the chromosome numbers of Al-chemilla species that belong to ser. Sericea and ser. Pubescentes, distributed in northeast Anatolia were reported (Hayırlıoâlu & Beyazoâlu 1997a).
The aim of this study was to investigate the chro-mosome numbers of 17 species of ser. Elatae
Rothm. In the ﬂora of Turkey, ser. Elatae is rep-resented by 24 species; the remaining 7 species (A. amonea, A. armeniaca, A. buseriana, A. hesii, A. holocycla, A. porrectidens and A. sciadiophylla) are not known from northeast Anatolia.
MATERIAL AND METHODS
The Alchemilla species of ser. Elatae used in this study were collected in the mountains of north-east Anatolia (Turkey) in July and August 1994. Voucher specimens are deposited in KTÜ. The studied specimens of each species are cited in “Results” below.
Actively growing root tips were used for the chro-mosome counts. The roots were cleaned of soil particles, then the root tips were cut off and pre-treated with 0.5% colchicine for 3 h (Beyazoâlu
et al. 1994), and then ﬁxed in an ethanol-acetic acid (3:1) solution for at least 24 h at 4°C. The root tips were hydrolyzed in 1 N HCl at 60°C for 15 min and then rinsed with tap water for a mini-mum of 2–3 min. Staining was carried out in Feul-gen for 1.5 h. Squashing was done in 45% acetic acid and the the preparations were mounted in Entellan.
In view of the serious difﬁculties in karyolog-ical analysis of Alchemilla mentioned above, the ﬁxation of root tips was, as a rule, repeated sev-eral times for each specimen studied. Of each
pop-ulation, 20–25 specimens for each species were collected and of these more than 15 permanent slides were prepared. The well-spread 10 meta-phase plates about 10 permanent slides were pho-tographed with an Olympus BH-2 camera and drawn from permanent slides deposited at the De-partment of Biology, Karadeniz Technical Uni-versity, Trabzon.
Asterisks (*) denote species for which chromo-some numbers have not been counted prior to this report. Numbers in parentheses indicate numbers of cells with chromosome numbers as indicated (e.g., “66–75 (2)” means that 2 cells were observed to have 66–75 chromosomes).
*Alchemilla barbatiflora Juz.
This is an Euxine element that grows on steep slopes at 1 700–2 400 m. It has been reported from Trabzon (Zigana Daâı at 1 700 m and from N side
of Soâanlı Daâı, above Çaykara at 2 000–2 200 m)
by Pawlowski and Walters (1972).
2n = 109–118(Fig. 1). Specimens: Rize, Ayder, Çamlı-hemñin, 1 450 m, 5.VII.1994, S. Hayırlıoâlu-Ayaz114: 109 (1), 109–113 (4); Trabzon, Araklı, Dagbañı, Kirazlı yay, 1 650 m, 9.VIII.1995, S. Hayırlıoâlu-Ayaz219: 109–113 (1), 112–118 (3), 113 (1).
*Alchemilla bornmuelleri Rothm.
This is an endemic and Iran-Turan element that grows on stream side banks at 1 200–1 800 m. It has not previously been recorded for northeast Anatolia.
2n = 64–75(Fig. 2). Specimens: Trabzon, Zigana Pass (Tunnel), 1 800 m, 17.VII.1995, S. Hayırlıoâlu-Ayaz 189: 64 (6), 66–75 (2), 70–72 (4), 72 (2), 75 (1).
*Alchemilla bursensis B. Pawl.
This is an endemic and Euxine element that grows in bogs under Fagus at 1 200–1 400 m. It has not
previously been recorded for northeast Anatolia.
2n = 96–102(Fig. 3). Specimens: Trabzon, Sultanmurat yay, Çaykara, 1 700 m, 30.VI.1993, S. Hayırlıoâlu-Ayaz1: 96 (4), 96–97 (2), 100–101 (2), 100–102 (2), 102 (2).
*Alchemilla ciminensis B. Pawl.
An endemic and Iran-Turan element that grows in Fagus and Picea forests and by streams at 2 200– 2 400 m. It has not previously been recorded for northeast Anatolia.
2n = 70–72(Fig. 4). Specimens: Çoruh, Ñavval Tepe
above Murgul, 2 200 m, 16.VII.1993, S. Hayırlıoâlu-Ayaz
12: 70 (1), 70–72 (4), 72 (2), 72–76 (5).
*Alchemilla erzincanensis B. Pawl.
An endemic and Iran-Turan element that grows along streams at 2 450 m. It has not previously been recorded for northeast Anatolia.
2n = 96–108(Fig. 5). Specimens: Trabzon, Zigana Pass, 1 800 m, 19.VIII.1993, S. Hayırlıoâlu-Ayaz 66: 96 (2), 96– 100 (3), 101–103 (1), 101–106 (2), 101–108 (3), 106 (2), 108 (1).
*Alchemilla hirsutiflora Rothm.
An endemic and Euxine element that grows on wet meadows and streamside banks at 1 800 m. It has not previously been recorded for northeast Anatolia.
2n = 90–102(Fig. 6). Specimens: Trabzon, Çaykara, Akdoâan köyü, 1 800 m, 21.VIII.1993, S. Hayırlıoâlu-Ayaz
95: 90 (1), 92 (4), 92–102 (1), 96–100 (2), 100–102 (3).
*Alchemilla hirtipedicellata B. Pawl.
An Euxine element that grows on meadows of forest slopes and wet meadows at 1 100–1 350 m. It was reported from Trabzon (Fol Köy at 1 100 m) by Pawlowski and Walters (1972).
2n = 86–96(Fig. 7). Specimens: Rize, Çayeli, Kaptan-paña, Çataldere Köyü, 1 350 m, 27.VII.1996, S. Hayırlıoâ
lu-Ayaz 308: 86 (2), 93–96 (4), 96 (6).
Alchemilla mollis Rothm.
This species grows by streams in Abies and Fagus
forests at 900–2 100 m. It has not previously been recorded for northeast Anatolia.
2n = 87–102(Fig. 8). Specimens: Gümüñhane, Gümü-ñhane to Trabzon, 1 500 m, 15.VIII.1993, S. Hayırlıoâ
lu-Ayaz 34: 88–90 (2), 90–92 (2), 91 (2), 102 (2); Rize, Anzer,
2 200 m, 21.VIII.1995, S. Hayırlıoâlu-Ayaz 212: 87–88 (2), 90–92 (2), 102 (3). Wegener (1967) reported 2n = 102– 106.
*Alchemilla orduensis B. Pawl.
An endemic and Euxine element that grows on stony mountain slopes at 1 800–2 200 m. It has not previously been recorded for northeast Ana-tolia.
2n = 98–108(Fig.9). Specimens: Trabzon, Zigana Daâı,
on the road of Kadırga, 2 200 m, 11.VIII.1993, S.
Hayırlıo-âlu-Ayaz 22: 98 (3), 98–100 (4), 106 (1), 106–108 (2), 108
*Alchemilla oriturcica Pawl.
An endemic and Iran-Turan element that grows on steep rocky slopes at 2 000–2 400 m. It was reported from Trabzon (Soâanlı daâı, above
Çay-kara at 2 000–2 200 m) by Pawlowski and Walters (1972).
2n = 86–100(Fig. 10). Specimens: Trabzon, Çaykara, Akdoâan Köyü, 1 400 m, 25.VIII.1993, S. Hayırlıoâlu-Ayaz
97: 86–90 (4), 90 (3), 104–106 (1), 106 (2).
*Alchemilla orthotricha Rothm.
An Euxine element that grows on meadows and streamside banks at 1 700–1 800 m. It was re-ported from Gümüñhane (Karagöl Daâı) and Trab-zon (Zigana Daâı above Hamsiköy ) by
Pawlow-ski and Walters (1972).
2n = 64 (Fig. 11). Specimens: Trabzon, Zigana Daâı,
above Hamsiköy, 1 800 m, 10.VII.1994, S. Hayırlıoâ
Figs. 1–8 (above and opposite). Mitotic metaphases in Alchemilla. The photographs (a) were taken with a BH 2 Olympus microscope. The drawings (b) were made with the aid of a drawing attachment connected to a BH 2 Olympus microscope. — 1: A. barbatiflora, 2n = 113. — 2: A. bornmuelleri, 2n = 75. — 3: A. bursensis, 2n = 102. — 4: A. ciminensis, 2n = 72. — 5: A. erzincanensis, 2n = 108. — 6: A. hirsutiflora, 2n = 100. — 7: A. hirtipedicellata, 2n = 96. — 8: A. mollis, 2n = 91. Scale bar = 10 µm.
Figs. 9–12. Mitotic metaphases in Alchemilla. The photographs (a) were taken with a BH 2 Olympus micro-scope. The drawings (b) were made with the aid of a drawing attachment connected to a BH 2 Olympus microscope. — 9: A. orduensis, 2n = 108. — 10: A. oriturcica, 2n = 90. — 11: A. orhotricha, 2n = 64. — 12: A. oxysepala, 2n = 107. Scale bar = 10 µm.
Alchemilla oxysepala Juz.
An Euxine element that grows on mountain mead-ows, forest and streamside banks at 1 600–1 800 m.
In the ﬂora of Turkey, this species is doubtfully recorded in northeast Anatolia (Pawlowski & Wal-ters 1972).
Ispir, 1 600 m, 10.VII.1995, S.Hayırlıoâlu-Ayaz 216: 95 (2), 103–104 (3), 104–107 (1), 107 (4). Ehrenberg (1945) reported 2n = 100; Turesson (1957) reported 2n = 105–109.
Alchemilla persica Rothm.
An Iran-Turan element that grows by streams at 1 850–2 800 m. It has not previously been re-corded for Northeast Anatolia.
2n = 96–106(Fig. 13). Specimens: Trabzon, Zigana Pass, 1 800 m, 20.VII.1993, S. Hayırlıoâlu-Ayaz 15: 96 (2), 96–104 (1), 104 (2); Trabzon, Araklı, Daâbañı, Kirazlı yay.,
1 650 m, 20.VII.1996, S. Hayırlıoâlu-Ayaz 300: 96–104 (2), 102 (1), 106 (4). Wegener (1967) reported 2n = 101– 106.
*Alchemilla sintenisii Rothm.
An endemic and Euxine element that grows in Alpine meadows at 1 800 m. It was recorded from Gümüñhane (Karagöl Daâı) by Pawlowski and
2n = 94–108(Fig. 14). Specimens: Trabzon, Zigana Tunnel and its around, 1 800 m, 7.VII.1995, S. Hayırlıoâ
lu-Ayaz 199: 94 (2), 94–96 (2), 94–98 (3), 106 (3).
*Alchemilla stricta Rothm.
This may be a Caucasian element that grows on marshy ground by lakes and streams and in Pinus
forest at 1 400–2 300 m. Alchemilla undecimloba
Juz. may be identical with this species according to the ﬂora of Turkey (Pawlowski & Walters 1972). It has not previously been recorded for northeast Anatolia.
2n = 66–100 (Fig. 15). Specimens: Rize, Cimil, Bañköy,
2 300 m, 11.VIII.1995, S. Hayırlıoâlu-Ayaz 302: 82 (2), 82–86 (1), 90–98 (2), 100 (3); Trabzon, Çaykara, Sultan-murat yay., 2 100 m, 11.VII.1993, S. Hayırlıoâlu-Ayaz 8: 66 (3), 67–68 (2), 70–75 (2), 82 (2), 97–98 (1).
*Alchemilla tiryalensis B.Pawl.
An endemic and Euxine element that grows on rocky igneous slopes at 2 300–2 700 m. It has been reported from Çoruh Tiryal Daâı, above Murgul,
2 300 m and Ñavval Tepe above Murgul, 2 700 m,
by Pawlowski and Walters (1972).
2n = 97–123 (Fig. 16). Specimens: Çoruh, Tiryal Daâı
above Murgul, 2 500 m, 4.VIII.1996, S. Hayırlıoâlu-Ayaz
303: 97 (3), 100 (1), 100–107 (2), 107 (2), 114–119 (1), 119–123 (3).
*Alchemilla ziganadagensis B. Pawl.
An endemic and Euxine element that grows in
Picea orientalis forests and woodlands at 1 700– 1 750 m. It was recorded from Trabzon (Zigana Daâı supra Hamsiköy) by Pawlowski and Walters
2n = 72(Fig. 17). Specimens: Trabzon, Zigana Daâı,
1 700 m, 19.VII.1994, S. Hayırlıoâlu-Ayaz 83: 72 (11).
Former karyological studies of Alchemilla spe-cies belonging to sect. Alchemilla have revealed that they are represented by a series of very high polyploid cytotypes with the chromosome num-bers ranging from 64 to ca. 224. About 75% of the species have chromosome numbers of 2n = 96–110 (Turesson 1957, Wegener 1967, Izmailow 1981).
Karyological data of ser. Elatae are still scarce. Some species were previously studied by Wegener (1967) from the Alps, Caucacus and a Botanical Garden in Austria, and by Ehrenberg (1945) and Turesson (1957) from the Alps. Wegener (1967) reported the chromosome numbers of A. mollis
(2n = 102–106) and A. persica (2n = 101–106) of ser. Elatae. Ehrenberg (1945) was the ﬁrst to re-port the chromosome number of A. oxysepala
(2n = 100). Later, Turesson (1957) reported the chromosome number of A. oxysepala as2n = 105– 109.
Comparison of reports by Wegener (1967), Ehrenberg (1945) and Turesson (1957) with the results of the present study shows that specimens collected from the Alps and Caucasus and from northeast Anatolia have different chromosome numbers. The diverging karyological results might result from intraspeciﬁc karyological differentia-tion. Such a differentiation has been found in
Al-Figs. 13–16. Mitotic metaphases in Alchemilla. The photographs (a) were taken with a BH 2 Olympus micro-scope. The drawings (b) were made with the aid of a drawing attachment connected to a BH 2 Olympus microscope. — 13: A. persica, 2n = 104. — 14: A. sintenisii, 2n = 106. — 15: A. stricta, 2n = 100. — 16: A. tiryalensis, 2n = 97. Scale bar = 10 µm.
to Wegener 1967). Similarly, Izmailow (1982) reported that specimens collected from various geographical regions have different chromosome
chemilla alpina (2n = 137–144 in Scandinavia, 2n = 119–122 in the Alps according to Turesson 1958; 2n = 119–129 in the central Alps according
Fig. 17. Mitotic
meta-phases in Alchemilla ziga-nadagensis, 2n = 72. Scale bar = 10 µm. The photograph (a) was taken with a BH 2 Olympus mi-croscope. The drawing (b) was made with the aid of a drawing attachment con-nected to a BH 2 Olympus microscope.
The investigations of Turesson (1957) and We-gener (1967) showed that that 75% of species of sect. Brevicaulon have 2n = 102–106. Informa-tion of genome, chromosome structure and karyo-gram are necessary in understanding the relation-ships of Alchemilla species. A taxonomic classiﬁ-cation of certain species of Alchemilla based on their chromosome numbers results in 4 main groups (Wegener 1967). In the ﬁrst group, the chromosome number is not more than 2n = 110, and in the second group it is 2n = 119–132. In the third and fourth groups, the chromosome num-bers are higher than in the other two groups, which clearly makes a difference between two groups (joining 1 and 2, and 3 and 4). The species stud-ied from northeast Anatolia belonging to the ser.
Sericeae, Pubescentes and Elatae (in the present study) represent the ﬁrst group.
Different series with the same chromosome numbers can occur in a single group. Thus, series are classiﬁed according to their morphological characters. Although the species are classiﬁed by a few certain morphological characters, a group may not be well distinguished morphologically.
Alchemillaorthotricha is similar to A. ziganada-gensis according to ﬂora of Turkey (Pawlowski & Walters 1972). Both species are distinguished by a few morphological features. Our results showed that A. orthotricha and A. ziganadagensis
have the chromosome numbers 2n = 64 and 2n = 72, respectively. We suggest that these numbers can be used as additional data to support morpho-logical separation of the species. Although the results of cytological studies have not been very important in the taxonomy of Alchemilla,
cyto-logical data can have taxonomic value, as this ex-ample shows.
In connection with diffuculties in an exact de-termination of chromosome numbers, the basic number as well as the degree of polyploidy in Al-chemilla remain problems. Gentscheff and Gus-tafsson (1940), as well as Gudjonsson (1941), sug-gested x = 7 is the basic number in Rosoideae. Later, Löve and Löve (1948) and Raven (1975) have suggested the basic number in Alchemilla is x = 8. The occurrence of the same basic number in the related genus Aphanes as well as the chro-mosome number 2n = 64 reported for some Al-chemilla species are in favour of ther latter opin-ion. Nevertheless, knowledge of the cytology of
Alchemilla is yet too insufﬁcient to determine the basic number with certainty (Wegener 1967).
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